Subphyla incertae sedis
A FUNGUS (/ˈfʌŋɡəs/ ; plural : FUNGI or FUNGUSES ) is any
member of the group of eukaryotic organisms that includes
microorganisms such as yeasts and molds , as well as the more familiar
mushrooms . These organisms are classified as a kingdom , FUNGI, which
is separate from the other eukaryotic life kingdoms of plants and
A characteristic that places fungi in a different kingdom from
plants, bacteria, and some protists is chitin in their cell walls .
Similar to animals, fungi are heterotrophs ; they acquire their food
by absorbing dissolved molecules, typically by secreting digestive
enzymes into their environment. Fungi do not photosynthesise . Growth
is their means of mobility, except for spores (a few of which are
flagellated), which may travel through the air or water. Fungi are the
principal decomposers in ecological systems. These and other
differences place fungi in a single group of related organisms, named
Eumycota (true fungi or Eumycetes), which share a common ancestor
(form a monophyletic group), an interpretation that is also strongly
supported by molecular phylogenetics . This fungal group is distinct
from the structurally similar myxomycetes (slime molds) and oomycetes
(water molds). The discipline of biology devoted to the study of fungi
is known as mycology (from the Greek μύκης mykes, meaning
"fungus"). In the past, mycology was regarded as a branch of botany ,
although it is now known fungi are genetically more closely related to
animals than to plants.
Abundant worldwide, most fungi are inconspicuous because of the small
size of their structures, and their cryptic lifestyles in soil or on
dead matter. Fungi include symbionts of plants, animals, or other
fungi and also parasites . They may become noticeable when fruiting ,
either as mushrooms or as molds. Fungi perform an essential role in
the decomposition of organic matter and have fundamental roles in
nutrient cycling and exchange in the environment. They have long been
used as a direct source of human food, in the form of mushrooms and
truffles ; as a leavening agent for bread; and in the fermentation of
various food products, such as wine , beer , and soy sauce . Since the
1940s, fungi have been used for the production of antibiotics , and,
more recently, various enzymes produced by fungi are used industrially
and in detergents . Fungi are also used as biological pesticides to
control weeds, plant diseases and insect pests. Many species produce
bioactive compounds called mycotoxins , such as alkaloids and
polyketides , that are toxic to animals including humans. The fruiting
structures of a few species contain psychotropic compounds and are
consumed recreationally or in traditional spiritual ceremonies . Fungi
can break down manufactured materials and buildings, and become
significant pathogens of humans and other animals. Losses of crops due
to fungal diseases (e.g., rice blast disease ) or food spoilage can
have a large impact on human food supplies and local economies.
The fungus kingdom encompasses an enormous diversity of taxa with
varied ecologies, life cycle strategies, and morphologies ranging from
unicellular aquatic chytrids to large mushrooms. However, little is
known of the true biodiversity of Kingdom Fungi, which has been
estimated at 1.5 million to 5 million species, with about 5% of these
having been formally classified. Ever since the pioneering 18th and
19th century taxonomical works of
Carl Linnaeus , Christian Hendrik
Persoon , and
Elias Magnus Fries , fungi have been classified
according to their morphology (e.g., characteristics such as spore
color or microscopic features) or physiology . Advances in molecular
genetics have opened the way for DNA analysis to be incorporated into
taxonomy, which has sometimes challenged the historical groupings
based on morphology and other traits.
Phylogenetic studies published
in the last decade have helped reshape the classification within
Kingdom Fungi, which is divided into one subkingdom , seven phyla ,
and ten subphyla.
* 1 Etymology
* 2 Characteristics
* 3 Diversity
* 4.1 History
* 5 Morphology
* 5.1 Microscopic structures
* 6 Growth and physiology
* 7.4 Other sexual processes
* 9 Taxonomy
* 9.1 Taxonomic groups
* 9.2 Fungus-like organisms
* 10.1.1 With plants
* 10.1.2 With algae and cyanobacteria
* 10.1.3 With insects
* 10.1.4 As pathogens and parasites
* 10.1.5 As targets of mycoparasites
* 12 Pathogenic mechanisms
* 13 Human use
* 13.1 Therapeutic uses
* 13.1.1 Modern chemotherapeutics
* 13.1.2 Traditional and folk medicine
* 13.2 Cultured foods
* 13.3 Edible and poisonous species
* 13.4 Pest control
* 13.7 Others
* 14 See also
* 15 References
* 15.1 Cited literature
* 16 External links
The English word fungus is directly adopted from the
(mushroom), used in the writings of
Horace and Pliny . This in turn
is derived from the Greek word sphongos (σφογγος "sponge"),
which refers to the macroscopic structures and morphology of mushrooms
and molds; the root is also used in other languages, such as the
German Schwamm ("sponge") and Schimmel ("mold"). The use of the word
mycology, which is derived from the Greek mykes (μύκης
"mushroom") and logos (λόγος "discourse"), to denote the
scientific study of fungi is thought to have originated in 1836 with
Miles Joseph Berkeley 's publication The English
Flora of Sir James Edward Smith, Vol. 5. A group of all the fungi
present in a particular area or geographic region is known as
mycobiota (plural noun, no singular), e.g., "the mycobiota of
FUNGAL HYPHAE CELLS
* Hyphal wall
Before the introduction of molecular methods for phylogenetic
analysis, taxonomists considered fungi to be members of the plant
kingdom because of similarities in lifestyle: both fungi and plants
are mainly immobile , and have similarities in general morphology and
growth habitat. Like plants, fungi often grow in soil and, in the case
of mushrooms , form conspicuous fruit bodies , which sometimes
resemble plants such as mosses . The fungi are now considered a
separate kingdom, distinct from both plants and animals, from which
they appear to have diverged around one billion years ago. Some
morphological, biochemical, and genetic features are shared with other
organisms, while others are unique to the fungi, clearly separating
them from the other kingdoms:
* With other eukaryotes : Fungal cells contain membrane-bound nuclei
with chromosomes that contain DNA with noncoding regions called
introns and coding regions called exons . Fungi have membrane-bound
cytoplasmic organelles such as mitochondria , sterol -containing
membranes, and ribosomes of the
80S type. They have a characteristic
range of soluble carbohydrates and storage compounds, including sugar
alcohols (e.g., mannitol ), disaccharides , (e.g., trehalose ), and
polysaccharides (e.g., glycogen , which is also found in animals ).
* With animals: Fungi lack chloroplasts and are heterotrophic
organisms and so require preformed organic compounds as energy
* With plants: Fungi have a cell wall and vacuoles . They
reproduce by both sexual and asexual means, and like basal plant
groups (such as ferns and mosses ) produce spores . Similar to mosses
and algae, fungi typically have haploid nuclei.
* With euglenoids and bacteria: Higher fungi, euglenoids, and some
bacteria produce the amino acid L-lysine in specific biosynthesis
steps, called the α-aminoadipate pathway .
* The cells of most fungi grow as tubular, elongated, and
thread-like (filamentous) structures called hyphae , which may contain
multiple nuclei and extend by growing at their tips. Each tip contains
a set of aggregated vesicles —cellular structures consisting of
proteins , lipids , and other organic molecules—called the
Spitzenkörper . Both fungi and oomycetes grow as filamentous hyphal
cells. In contrast, similar-looking organisms, such as filamentous
green algae , grow by repeated cell division within a chain of cells.
There are also single-celled fungi (yeasts ) that do not form hyphae,
and some fungi have both hyphal and yeast forms.
* In common with some plant and animal species, more than 70 fungal
species display bioluminescence .
* Some species grow as unicellular yeasts that reproduce by budding
or binary fission .
Dimorphic fungi can switch between a yeast phase
and a hyphal phase in response to environmental conditions.
* The fungal cell wall is composed of glucans and chitin ; while
glucans are also found in plants and chitin in the exoskeleton of
arthropods , fungi are the only organisms that combine these two
structural molecules in their cell wall. Unlike those of plants and
oomycetes, fungal cell walls do not contain cellulose.
Omphalotus nidiformis , a bioluminescent mushroom
Most fungi lack an efficient system for the long-distance transport
of water and nutrients, such as the xylem and phloem in many plants.
To overcome this limitation, some fungi, such as
Armillaria , form
rhizomorphs , which resemble and perform functions similar to the
roots of plants. As eukaryotes, fungi possess a biosynthetic pathway
for producing terpenes that uses mevalonic acid and pyrophosphate as
chemical building blocks .
Plants and some other organisms have an
additional terpene biosynthesis pathway in their chloroplasts, a
structure fungi and animals do not have. Fungi produce several
secondary metabolites that are similar or identical in structure to
those made by plants. Many of the plant and fungal enzymes that make
these compounds differ from each other in sequence and other
characteristics, which indicates separate origins and convergent
evolution of these enzymes in the fungi and plants.
Bracket fungi on a tree stump
Fungi have a worldwide distribution, and grow in a wide range of
habitats, including extreme environments such as deserts or areas with
high salt concentrations or ionizing radiation , as well as in deep
sea sediments. Some can survive the intense UV and cosmic radiation
encountered during space travel. Most grow in terrestrial
environments, though several species live partly or solely in aquatic
habitats, such as the chytrid fungus
Batrachochytrium dendrobatidis ,
a parasite that has been responsible for a worldwide decline in
amphibian populations. This organism spends part of its life cycle as
a motile zoospore , enabling it to propel itself through water and
enter its amphibian host. Other examples of aquatic fungi include
those living in hydrothermal areas of the ocean.
Around 100,000 species of fungi have been formally described by
taxonomists , but the global biodiversity of the fungus kingdom is
not fully understood. On the basis of observations of the ratio of
the number of fungal species to the number of plant species in
selected environments, the fungal kingdom has been estimated to
contain about 1.5 million species. A recent (2011) estimate suggests
there may be over 5 million species. In mycology, species have
historically been distinguished by a variety of methods and concepts.
Classification based on morphological characteristics, such as the
size and shape of spores or fruiting structures, has traditionally
dominated fungal taxonomy. Species may also be distinguished by their
biochemical and physiological characteristics, such as their ability
to metabolize certain biochemicals, or their reaction to chemical
tests . The biological species concept discriminates species based on
their ability to mate . The application of molecular tools, such as
DNA sequencing and phylogenetic analysis, to study diversity has
greatly enhanced the resolution and added robustness to estimates of
genetic diversity within various taxonomic groups. Two types of
Mycology is the branch of biology concerned with the systematic study
of fungi, including their genetic and biochemical properties, their
taxonomy, and their use to humans as a source of medicine, food, and
psychotropic substances consumed for religious purposes, as well as
their dangers, such as poisoning or infection. The field of
phytopathology , the study of plant diseases, is closely related
because many plant pathogens are fungi. In 1729, Pier A. Micheli
first published descriptions of fungi.
The use of fungi by humans dates back to prehistory; Ötzi the Iceman
, a well-preserved mummy of a 5,300-year-old
Neolithic man found
frozen in the Austrian Alps, carried two species of polypore mushrooms
that may have been used as tinder (
Fomes fomentarius ), or for
medicinal purposes (
Piptoporus betulinus ). Ancient peoples have used
fungi as food sources–often unknowingly–for millennia, in the
preparation of leavened bread and fermented juices. Some of the oldest
written records contain references to the destruction of crops that
were probably caused by pathogenic fungi.
Mycology is a relatively new science that became systematic after the
development of the microscope in the 17th century. Although fungal
spores were first observed by
Giambattista della Porta in 1588, the
seminal work in the development of mycology is considered to be the
Pier Antonio Micheli
Pier Antonio Micheli 's 1729 work Nova plantarum
genera. Micheli not only observed spores but also showed that, under
the proper conditions, they could be induced into growing into the
same species of fungi from which they originated. Extending the use
of the binomial system of nomenclature introduced by
Carl Linnaeus in
Species plantarum (1753), the Dutch Christian Hendrik Persoon
(1761–1836) established the first classification of mushrooms with
such skill so as to be considered a founder of modern mycology. Later,
Elias Magnus Fries (1794–1878) further elaborated the classification
of fungi, using spore color and various microscopic characteristics,
methods still used by taxonomists today. Other notable early
contributors to mycology in the 17th–19th and early 20th centuries
Miles Joseph Berkeley ,
August Carl Joseph Corda , Anton de
Bary , the brothers Louis René and
Charles Tulasne , Arthur H. R.
Buller , Curtis G. Lloyd , and
Pier Andrea Saccardo . The 20th century
has seen a modernization of mycology that has come from advances in
biochemistry , genetics , molecular biology , and biotechnology . The
DNA sequencing technologies and phylogenetic analysis has
provided new insights into fungal relationships and biodiversity , and
has challenged traditional morphology-based groupings in fungal
An environmental isolate of
Most fungi grow as hyphae , which are cylindrical, thread-like
structures 2–10 µm in diameter and up to several centimeters in
Hyphae grow at their tips (apices); new hyphae are typically
formed by emergence of new tips along existing hyphae by a process
called branching, or occasionally growing hyphal tips fork, giving
rise to two parallel-growing hyphae.
Hyphae also sometimes fuse when
they come into contact, a process called hyphal fusion (or anastamosis
). These growth processes lead to the development of a mycelium , an
interconnected network of hyphae.
Hyphae can be either septate or
coenocytic . Septate hyphae are divided into compartments separated by
cross walls (internal cell walls, called septa, that are formed at
right angles to the cell wall giving the hypha its shape), with each
compartment containing one or more nuclei; coenocytic hyphae are not
compartmentalized. Septa have pores that allow cytoplasm , organelles
, and sometimes nuclei to pass through; an example is the dolipore
septum in fungi of the phylum Basidiomycota.
Coenocytic hyphae are in
essence multinucleate supercells.
Many species have developed specialized hyphal structures for
nutrient uptake from living hosts; examples include haustoria in
plant-parasitic species of most fungal phyla, and arbuscules of
several mycorrhizal fungi, which penetrate into the host cells to
Although fungi are opisthokonts —a grouping of evolutionarily
related organisms broadly characterized by a single posterior
flagellum —all phyla except for the chytrids have lost their
posterior flagella. Fungi are unusual among the eukaryotes in having
a cell wall that, in addition to glucans (e.g., β-1,3-glucan) and
other typical components, also contains the biopolymer chitin.
Fungal mycelia can become visible to the naked eye, for example, on
various surfaces and substrates , such as damp walls and spoiled food,
where they are commonly called molds . Mycelia grown on solid agar
media in laboratory petri dishes are usually referred to as colonies .
These colonies can exhibit growth shapes and colors (due to spores or
pigmentation ) that can be used as diagnostic features in the
identification of species or groups. Some individual fungal colonies
can reach extraordinary dimensions and ages as in the case of a clonal
Armillaria solidipes , which extends over an area of more
than 900 ha (3.5 square miles), with an estimated age of nearly 9,000
The apothecium —a specialized structure important in sexual
reproduction in the ascomycetes—is a cup-shaped fruit body that is
often macroscopic and holds the hymenium , a layer of tissue
containing the spore-bearing cells. The fruit bodies of the
basidiomycetes (basidiocarps ) and some ascomycetes can sometimes grow
very large, and many are well known as mushrooms .
GROWTH AND PHYSIOLOGY
Mold growth covering a decaying peach . The frames were taken
approximately 12 hours apart over a period of six days.
The growth of fungi as hyphae on or in solid substrates or as single
cells in aquatic environments is adapted for the efficient extraction
of nutrients, because these growth forms have high surface area to
volume ratios .
Hyphae are specifically adapted for growth on solid
surfaces, and to invade substrates and tissues. They can exert large
penetrative mechanical forces; for example, many plant pathogens ,
Magnaporthe grisea , form a structure called an appressorium
that evolved to puncture plant tissues. The pressure generated by the
appressorium, directed against the plant epidermis , can exceed 8
megapascals (1,200 psi). The filamentous fungus Paecilomyces
lilacinus uses a similar structure to penetrate the eggs of nematodes
The mechanical pressure exerted by the appressorium is generated from
physiological processes that increase intracellular turgor by
producing osmolytes such as glycerol . Adaptations such as these are
complemented by hydrolytic enzymes secreted into the environment to
digest large organic molecules—such as polysaccharides , proteins ,
and lipids —into smaller molecules that may then be absorbed as
nutrients. The vast majority of filamentous fungi grow in a polar
fashion (extending in one direction) by elongation at the tip (apex)
of the hypha. Other forms of fungal growth include intercalary
extension (longitudinal expansion of hyphal compartments that are
below the apex) as in the case of some endophytic fungi, or growth by
volume expansion during the development of mushroom stipes and other
large organs. Growth of fungi as multicellular structures consisting
of somatic and reproductive cells—a feature independently evolved in
animals and plants —has several functions, including the development
of fruit bodies for dissemination of sexual spores (see above) and
biofilms for substrate colonization and intercellular communication .
The fungi are traditionally considered heterotrophs , organisms that
rely solely on carbon fixed by other organisms for metabolism . Fungi
have evolved a high degree of metabolic versatility that allows them
to use a diverse range of organic substrates for growth, including
simple compounds such as nitrate , ammonia , acetate , or ethanol .
In some species the pigment melanin may play a role in extracting
energy from ionizing radiation , such as gamma radiation . This form
of "radiotrophic" growth has been described for only a few species,
the effects on growth rates are small, and the underlying biophysical
and biochemical processes are not well known. This process might bear
similarity to CO2 fixation via visible light , but instead uses
ionizing radiation as a source of energy.
Fungal reproduction is complex, reflecting the differences in
lifestyles and genetic makeup within this diverse kingdom of
organisms. It is estimated that a third of all fungi reproduce using
more than one method of propagation; for example, reproduction may
occur in two well-differentiated stages within the life cycle of a
species, the teleomorph and the anamorph . Environmental conditions
trigger genetically determined developmental states that lead to the
creation of specialized structures for sexual or asexual reproduction.
These structures aid reproduction by efficiently dispersing spores or
spore-containing propagules .
Asexual reproduction occurs via vegetative spores (conidia ) or
through mycelial fragmentation . Mycelial fragmentation occurs when a
fungal mycelium separates into pieces, and each component grows into a
separate mycelium. Mycelial fragmentation and vegetative spores
maintain clonal populations adapted to a specific niche , and allow
more rapid dispersal than sexual reproduction. The "Fungi imperfecti"
(fungi lacking the perfect or sexual stage) or
all the species that lack an observable sexual cycle. Deuteromycota
is not an accepted taxonomic clade, and is now taken to mean simply
fungi that lack a known sexual stage.
Mating in fungi
Sexual reproduction with meiosis has been directly observed in all
fungal phyla except
Glomeromycota (genetic analysis suggests meiosis
Glomeromycota as well). It differs in many aspects from sexual
reproduction in animals or plants. Differences also exist between
fungal groups and can be used to discriminate species by morphological
differences in sexual structures and reproductive strategies. Mating
experiments between fungal isolates may identify species on the basis
of biological species concepts. The major fungal groupings have
initially been delineated based on the morphology of their sexual
structures and spores; for example, the spore-containing structures,
asci and basidia , can be used in the identification of ascomycetes
and basidiomycetes, respectively. Some species may allow mating only
between individuals of opposite mating type , whereas others can mate
and sexually reproduce with any other individual or itself. Species of
the former mating system are called heterothallic , and of the latter
Most fungi have both a haploid and a diploid stage in their life
cycles. In sexually reproducing fungi, compatible individuals may
combine by fusing their hyphae together into an interconnected
network; this process, anastomosis , is required for the initiation of
the sexual cycle. Many ascomycetes and basidiomycetes go through a
dikaryotic stage, in which the nuclei inherited from the two parents
do not combine immediately after cell fusion, but remain separate in
the hyphal cells (see heterokaryosis ). The 8-spore asci of
Morchella elata , viewed with phase contrast microscopy
In ascomycetes, dikaryotic hyphae of the hymenium (the spore-bearing
tissue layer) form a characteristic hook at the hyphal septum. During
cell division , formation of the hook ensures proper distribution of
the newly divided nuclei into the apical and basal hyphal
compartments. An ascus (plural asci) is then formed, in which
karyogamy (nuclear fusion) occurs. Asci are embedded in an ascocarp ,
or fruiting body.
Karyogamy in the asci is followed immediately by
meiosis and the production of ascospores . After dispersal, the
ascospores may germinate and form a new haploid mycelium.
Sexual reproduction in basidiomycetes is similar to that of the
ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic
mycelium. However, the dikaryotic phase is more extensive in the
basidiomycetes, often also present in the vegetatively growing
mycelium. A specialized anatomical structure, called a clamp
connection , is formed at each hyphal septum. As with the structurally
similar hook in the ascomycetes, the clamp connection in the
basidiomycetes is required for controlled transfer of nuclei during
cell division, to maintain the dikaryotic stage with two genetically
different nuclei in each hyphal compartment. A basidiocarp is formed
in which club-like structures known as basidia generate haploid
basidiospores after karyogamy and meiosis. The most commonly known
basidiocarps are mushrooms, but they may also take other forms (see
In glomeromycetes (formerly zygomycetes), haploid hyphae of two
individuals fuse, forming a gametangium , a specialized cell structure
that becomes a fertile gamete -producing cell. The gametangium
develops into a zygospore , a thick-walled spore formed by the union
of gametes. When the zygospore germinates, it undergoes meiosis ,
generating new haploid hyphae, which may then form asexual
sporangiospores . These sporangiospores allow the fungus to rapidly
disperse and germinate into new genetically identical haploid fungal
Both asexual and sexual spores or sporangiospores are often actively
dispersed by forcible ejection from their reproductive structures.
This ejection ensures exit of the spores from the reproductive
structures as well as traveling through the air over long distances.
The bird's nest fungus
Specialized mechanical and physiological mechanisms, as well as spore
surface structures (such as hydrophobins ), enable efficient spore
ejection. For example, the structure of the spore-bearing cells in
some ascomycete species is such that the buildup of substances
affecting cell volume and fluid balance enables the explosive
discharge of spores into the air. The forcible discharge of single
spores termed ballistospores involves formation of a small drop of
water (Buller's drop), which upon contact with the spore leads to its
projectile release with an initial acceleration of more than 10,000 g
; the net result is that the spore is ejected 0.01–0.02 cm,
sufficient distance for it to fall through the gills or pores into the
air below. Other fungi, like the puffballs , rely on alternative
mechanisms for spore release, such as external mechanical forces. The
bird\'s nest fungi use the force of falling water drops to liberate
the spores from cup-shaped fruiting bodies. Another strategy is seen
in the stinkhorns , a group of fungi with lively colors and putrid
odor that attract insects to disperse their spores.
OTHER SEXUAL PROCESSES
Besides regular sexual reproduction with meiosis, certain fungi, such
as those in the genera
Aspergillus , may exchange
genetic material via parasexual processes, initiated by anastomosis
between hyphae and plasmogamy of fungal cells. The frequency and
relative importance of parasexual events is unclear and may be lower
than other sexual processes. It is known to play a role in
intraspecific hybridization and is likely required for hybridization
between species, which has been associated with major events in fungal
Evolution of fungi
In contrast to plants and animals , the early fossil record of the
fungi is meager. Factors that likely contribute to the
under-representation of fungal species among fossils include the
nature of fungal fruiting bodies , which are soft, fleshy, and easily
degradable tissues and the microscopic dimensions of most fungal
structures, which therefore are not readily evident. Fungal fossils
are difficult to distinguish from those of other microbes, and are
most easily identified when they resemble extant fungi. Often
recovered from a permineralized plant or animal host, these samples
are typically studied by making thin-section preparations that can be
examined with light microscopy or transmission electron microscopy .
Researchers study compression fossils by dissolving the surrounding
matrix with acid and then using light or scanning electron microscopy
to examine surface details.
The earliest fossils possessing features typical of fungi date to the
Paleoproterozoic era, some 2,400 million years ago (Ma ); these
multicellular benthic organisms had filamentous structures capable of
anastomosis . Other studies (2009) estimate the arrival of fungal
organisms at about 760–1060 Ma on the basis of comparisons of the
rate of evolution in closely related groups. For much of the
Paleozoic Era (542–251 Ma), the fungi appear to have been aquatic
and consisted of organisms similar to the extant chytrids in having
flagellum-bearing spores. The evolutionary adaptation from an aquatic
to a terrestrial lifestyle necessitated a diversification of
ecological strategies for obtaining nutrients, including parasitism ,
saprobism , and the development of mutualistic relationships such as
mycorrhiza and lichenization. Recent (2009) studies suggest that the
ancestral ecological state of the
Ascomycota was saprobism, and that
independent lichenization events have occurred multiple times.
It is presumed that the fungi colonized the land during the Cambrian
(542–488.3 Ma), long before land plants. Fossilized hyphae and
spores recovered from the
Ordovician of Wisconsin (460 Ma) resemble
Glomerales , and existed at a time when the land flora
likely consisted of only non-vascular bryophyte -like plants.
Prototaxites , which was probably a fungus or lichen, would have been
the tallest organism of the late
Silurian . Fungal fossils do not
become common and uncontroversial until the early Devonian
(416–359.2 Ma), when they occur abundantly in the
Rhynie chert ,
Chytridiomycota . At about this same time,
approximately 400 Ma, the
Basidiomycota diverged, and
all modern classes of fungi were present by the Late Carboniferous
(Pennsylvanian , 318.1–299 Ma).
Lichen -like fossils have been found in the
Doushantuo Formation in
southern China dating back to 635–551 Ma.
Lichens formed a
component of the early terrestrial ecosystems, and the estimated age
of the oldest terrestrial lichen fossil is 400 Ma; this date
corresponds to the age of the oldest known sporocarp fossil, a
Paleopyrenomycites species found in the Rhynie Chert. The oldest
fossil with microscopic features resembling modern-day basidiomycetes
is Palaeoancistrus, found permineralized with a fern from the
Pennsylvanian. Rare in the fossil record are the Homobasidiomycetes
(a taxon roughly equivalent to the mushroom-producing species of the
Agaricomycetes ). Two amber -preserved specimens provide evidence that
the earliest known mushroom-forming fungi (the extinct species
Archaeomarasmius leggetti ) appeared during the late
Cretaceous , 90
Some time after the Permian–
Triassic extinction event (251.4 Ma), a
fungal spike (originally thought to be an extraordinary abundance of
fungal spores in sediments ) formed, suggesting that fungi were the
dominant life form at this time, representing nearly 100% of the
available fossil record for this period. However, the relative
proportion of fungal spores relative to spores formed by algal species
is difficult to assess, the spike did not appear worldwide, and in
many places it did not fall on the Permian–
Although commonly included in botany curricula and textbooks, fungi
are more closely related to animals than to plants and are placed with
the animals in the monophyletic group of opisthokonts . Analyses
using molecular phylogenetics support a monophyletic origin of the
Fungi. The taxonomy of the Fungi is in a state of constant flux,
especially due to recent research based on DNA comparisons. These
current phylogenetic analyses often overturn classifications based on
older and sometimes less discriminative methods based on morphological
features and biological species concepts obtained from experimental
There is no unique generally accepted system at the higher taxonomic
levels and there are frequent name changes at every level, from
species upwards. Efforts among researchers are now underway to
establish and encourage usage of a unified and more consistent
nomenclature . Fungal species can also have multiple scientific
names depending on their life cycle and mode (sexual or asexual) of
reproduction. Web sites such as
Index Fungorum and ITIS list current
names of fungal species (with cross-references to older synonyms).
The 2007 classification of Kingdom Fungi is the result of a
large-scale collaborative research effort involving dozens of
mycologists and other scientists working on fungal taxonomy. It
recognizes seven phyla , two of which—the
Ascomycota and the
Basidiomycota—are contained within a branch representing subkingdom
Dikarya, the most species rich and familiar group, including all the
mushrooms, most food-spoilage molds, most plant pathogenic fungi, and
the beer, wine, and bread yeasts. The accompanying cladogram depicts
the major fungal taxa and their relationship to opisthokont and
unikont organisms, based on the work of Philippe Silar and "The
Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for
Basic and Applied Research". The lengths of the branches are not
proportional to evolutionary distances.
List of fungal orders Main groups of fungi.
The major phyla (sometimes called divisions) of fungi have been
classified mainly on the basis of characteristics of their sexual
reproductive structures. Currently, seven phyla are proposed:
Microsporidia, Chytridiomycota, Blastocladiomycota,
Neocallimastigomycota, Glomeromycota, Ascomycota, and Basidiomycota.
Phylogenetic analysis has demonstrated that the
unicellular parasites of animals and protists, are fairly recent and
highly derived endobiotic fungi (living within the tissue of another
species). One 2006 study concludes that the
Microsporidia are a
sister group to the true fungi; that is, they are each other's closest
evolutionary relative. Hibbett and colleagues suggest that this
analysis does not clash with their classification of the Fungi, and
Microsporidia are elevated to phylum status, it is
acknowledged that further analysis is required to clarify evolutionary
relationships within this group.
Chytridiomycota are commonly known as chytrids. These fungi are
Chytrids and their close relatives
Blastocladiomycota (below) are the only
fungi with active motility, producing zoospores that are capable of
active movement through aqueous phases with a single flagellum ,
leading early taxonomists to classify them as protists . Molecular
phylogenies , inferred from rRNA sequences in ribosomes , suggest that
Chytrids are a basal group divergent from the other fungal phyla,
consisting of four major clades with suggestive evidence for paraphyly
or possibly polyphyly .
Blastocladiomycota were previously considered a taxonomic clade
within the Chytridiomycota. Recent molecular data and ultrastructural
characteristics, however, place the
Blastocladiomycota as a sister
clade to the Zygomycota, Glomeromycota, and
Basidiomycota). The blastocladiomycetes are saprotrophs , feeding on
decomposing organic matter, and they are parasites of all eukaryotic
groups. Unlike their close relatives, the chytrids, most of which
exhibit zygotic meiosis , the blastocladiomycetes undergo sporic
Neocallimastigomycota were earlier placed in the phylum
Chytridomycota. Members of this small phylum are anaerobic organisms ,
living in the digestive system of larger herbivorous mammals and in
other terrestrial and aquatic environments enriched in cellulose
(e.g., domestic waste landfill sites). They lack mitochondria but
contain hydrogenosomes of mitochondrial origin. As in the related
chrytrids, neocallimastigomycetes form zoospores that are posteriorly
uniflagellate or polyflagellate.
Arbuscular mycorrhiza seen
Flax root cortical cells containing paired
arbuscules. Diagram of an apothecium (the typical cup-like
reproductive structure of Ascomycetes) showing sterile tissues as well
as developing and mature asci.
Members of the
Glomeromycota form arbuscular mycorrhizae , a form of
mutualist symbiosis wherein fungal hyphae invade plant root cells and
both species benefit from the resulting increased supply of nutrients.
Glomeromycota species reproduce asexually. The symbiotic
association between the
Glomeromycota and plants is ancient, with
evidence dating to 400 million years ago. Formerly part of the
Zygomycota (commonly known as 'sugar' and 'pin' molds), the
Glomeromycota were elevated to phylum status in 2001 and now replace
the older phylum Zygomycota. Fungi that were placed in the Zygomycota
are now being reassigned to the Glomeromycota, or the subphyla
Kickxellomycotina , the
Zoopagomycotina and the
Entomophthoromycotina . Some well-known
examples of fungi formerly in the
Zygomycota include black bread mold
Rhizopus stolonifer ), and
Pilobolus species, capable of ejecting
spores several meters through the air. Medically relevant genera
Rhizomucor , and
Ascomycota , commonly known as sac fungi or ascomycetes,
constitute the largest taxonomic group within the Eumycota. These
fungi form meiotic spores called ascospores , which are enclosed in a
special sac-like structure called an ascus . This phylum includes
morels , a few mushrooms and truffles , unicellular yeasts (e.g., of
Pichia , and Candida ), and
many filamentous fungi living as saprotrophs, parasites, and
mutualistic symbionts (e.g. lichens). Prominent and important genera
of filamentous ascomycetes include
Fusarium , and
Claviceps . Many ascomycete species have only been
observed undergoing asexual reproduction (called anamorphic species),
but analysis of molecular data has often been able to identify their
closest teleomorphs in the Ascomycota. Because the products of
meiosis are retained within the sac-like ascus, ascomycetes have been
used for elucidating principles of genetics and heredity (e.g.,
Neurospora crassa ).
Members of the
Basidiomycota , commonly known as the club fungi or
basidiomycetes, produce meiospores called basidiospores on club-like
stalks called basidia . Most common mushrooms belong to this group, as
well as rust and smut fungi , which are major pathogens of grains.
Other important basidiomycetes include the maize pathogen Ustilago
maydis , human commensal species of the genus
Malassezia , and the
opportunistic human pathogen,
Cryptococcus neoformans .
Because of similarities in morphology and lifestyle, the slime molds
(mycetozoans , plasmodiophorids , acrasids ,
labyrinthulids , now in
Excavata , Opisthokonta
Stramenopiles , respectively), water molds (oomycetes ) and
Stramenopiles ) were formerly classified in the
kingdom Fungi, in groups like
Phycomycetes . The slime molds were studied also as protozoans ,
leading to a ambiregnal , duplicated taxonomy.
Unlike true fungi, the cell walls of oomycetes contain cellulose and
lack chitin . Hyphochytrids have both chitin and cellulose. Slime
molds lack a cell wall during the assimilative phase (except
labyrinthulids, which have a wall of scales), and ingest nutrients by
ingestion (phagocytosis , except labyrinthulids) rather than
absorption (osmotrophy , as fungi, labyrinthulids, oomycetes and
hyphochytrids). Neither water molds nor slime molds are closely
related to the true fungi, and, therefore, taxonomists no longer group
them in the kingdom Fungi. Nonetheless, studies of the oomycetes and
myxomycetes are still often included in mycology textbooks and primary
Amoebidiales are opisthokont protists , previously
thought to be zygomycete fungi. Other groups now in Opisthokonta
Ichthyosporea ) were also at given time
classified as fungi. The genus
Blastocystis , now in
was originally classified as a yeast.
Ellobiopsis , now in
was considered a chytrid. The bacteria were also included in fungi in
some classifications, as the group Schizomycetes.
Rozellida clade, including the "ex-chytrid"
Rozella , is a
genetically disparate group known mostly from environmental DNA
sequences that is a sister group to fungi. Members of the group that
have been isolated lack the chitinous cell wall that is characteristic
The nucleariids , protists currently grouped in the Choanozoa
Opisthokonta ), may be the next sister group to the eumycete clade,
and as such could be included in an expanded fungal kingdom.
Actinobacteria ), a group with many filamentous
bacteria, were also long believed to be fungi.
A pin mold decomposing a peach
Although often inconspicuous, fungi occur in every environment on
Earth and play very important roles in most ecosystems . Along with
bacteria, fungi are the major decomposers in most terrestrial (and
some aquatic) ecosystems, and therefore play a critical role in
biogeochemical cycles and in many food webs . As decomposers, they
play an essential role in nutrient cycling , especially as saprotrophs
and symbionts , degrading organic matter to inorganic molecules, which
can then re-enter anabolic metabolic pathways in plants or other
Many fungi have important symbiotic relationships with organisms from
most if not all Kingdoms . These interactions can be mutualistic or
antagonistic in nature, or in the case of commensal fungi are of no
apparent benefit or detriment to the host.
Mycorrhizal symbiosis between plants and fungi is one of the most
well-known plant–fungus associations and is of significant
importance for plant growth and persistence in many ecosystems; over
90% of all plant species engage in mycorrhizal relationships with
fungi and are dependent upon this relationship for survival. The
dark filaments are hyphae of the endophytic fungus Neotyphodium
coenophialum in the intercellular spaces of tall fescue leaf sheath
The mycorrhizal symbiosis is ancient, dating to at least 400 million
years ago. It often increases the plant's uptake of inorganic
compounds, such as nitrate and phosphate from soils having low
concentrations of these key plant nutrients. The fungal partners may
also mediate plant-to-plant transfer of carbohydrates and other
nutrients. Such mycorrhizal communities are called "common mycorrhizal
networks". A special case of mycorrhiza is myco-heterotrophy ,
whereby the plant parasitizes the fungus, obtaining all of its
nutrients from its fungal symbiont. Some fungal species inhabit the
tissues inside roots, stems, and leaves, in which case they are called
endophytes. Similar to mycorrhiza, endophytic colonization by fungi
may benefit both symbionts; for example, endophytes of grasses impart
to their host increased resistance to herbivores and other
environmental stresses and receive food and shelter from the plant in
Algae And Cyanobacteria
Lobaria pulmonaria , a symbiosis of fungal, algal ,
and cyanobacterial species
Lichens are a symbiotic relationship between fungi and photosynthetic
algae or cyanobacteria . The photosynthetic partner in the
relationship is referred to in lichen terminology as a "photobiont".
The fungal part of the relationship is composed mostly of various
species of ascomycetes and a few basidiomycetes .
Lichens occur in
every ecosystem on all continents, play a key role in soil formation
and the initiation of biological succession , and are prominent in
some extreme environments, including polar , alpine , and semiarid
desert regions. They are able to grow on inhospitable surfaces,
including bare soil, rocks, tree bark , wood, shells, barnacles and
leaves. As in mycorrhizas , the photobiont provides sugars and other
carbohydrates via photosynthesis to the fungus, while the fungus
provides minerals and water to the photobiont. The functions of both
symbiotic organisms are so closely intertwined that they function
almost as a single organism; in most cases the resulting organism
differs greatly from the individual components. Lichenization is a
common mode of nutrition for fungi; around 20% of fungi—between
17,500 and 20,000 described species—are lichenized. Characteristics
common to most lichens include obtaining organic carbon by
photosynthesis, slow growth, small size, long life, long-lasting
(seasonal) vegetative reproductive structures, mineral nutrition
obtained largely from airborne sources, and greater tolerance of
desiccation than most other photosynthetic organisms in the same
Many insects also engage in mutualistic relationships with fungi.
Several groups of ants cultivate fungi in the order
their primary food source, while ambrosia beetles cultivate various
species of fungi in the bark of trees that they infest. Likewise,
females of several wood wasp species (genus
Sirex ) inject their eggs
together with spores of the wood-rotting fungus Amylostereum areolatum
into the sapwood of pine trees; the growth of the fungus provides
ideal nutritional conditions for the development of the wasp larvae.
At least one species of stingless bee has a relationship with a fungus
in the genus
Monascus , where the larvae consume and depend on fungus
transferred from old to new nests.
Termites on the African savannah
are also known to cultivate fungi, and yeasts of the genera Candida
and Lachancea inhabit the gut of a wide range of insects, including
neuropterans , beetles , and cockroaches ; it is not known whether
these fungi benefit their hosts. Fungi ingrowing dead wood are
essential for xylophagous insects (e.g. woodboring beetles ). They
deliver nutrients needed by xylophages to nutritionally scarce dead
wood . Thanks to this nutritional enrichment the larvae of woodboring
insect is able to grow and develop to adulthood. The larvae of many
families of fungicolous flies, particularly those within the
Sciaroidea such as the
Mycetophilidae and some
Keroplatidae feed on fungal fruiting bodies and sterile mycorrhizae .
As Pathogens And Parasites
The plant pathogen Aecidium magellanicum causes calafate rust ,
seen here on a
Berberis shrub in Chile.
Many fungi are parasites on plants, animals (including humans), and
other fungi. Serious pathogens of many cultivated plants causing
extensive damage and losses to agriculture and forestry include the
rice blast fungus
Magnaporthe oryzae , tree pathogens such as
Ophiostoma ulmi and
Ophiostoma novo-ulmi causing
Dutch elm disease
Dutch elm disease
Cryphonectria parasitica responsible for chestnut blight , and
plant pathogens in the genera
Alternaria , and
Cochliobolus . Some carnivorous fungi , like
Paecilomyces lilacinus ,
are predators of nematodes , which they capture using an array of
specialized structures such as constricting rings or adhesive nets.
Some fungi can cause serious diseases in humans, several of which may
be fatal if untreated. These include aspergillosis , candidiasis ,
coccidioidomycosis , cryptococcosis , histoplasmosis , mycetomas , and
paracoccidioidomycosis . Furthermore, persons with immuno-deficiencies
are particularly susceptible to disease by genera such as Aspergillus
, Candida , Cryptoccocus ,
Histoplasma , and
Pneumocystis . Other
fungi can attack eyes, nails, hair, and especially skin, the so-called
dermatophytic and keratinophilic fungi, and cause local infections
such as ringworm and athlete\'s foot . Fungal spores are also a cause
of allergies , and fungi from different taxonomic groups can evoke
As Targets Of Mycoparasites
The organisms which parasitize fungi are known as mycoparasitic
organisms. Certain species of the
Pythium genus, which are oomycetes ,
have potential as biocontrol agents against certain fungi. Fungi can
also act as mycoparasites or antagonists of other fungi, such as
Hypomyces chrysospermus , which grows on bolete mushrooms.
Fungi can become the target of infection by mycoviruses .
Ergotamine , a major mycotoxin produced by
which if ingested can cause gangrene , convulsions, and hallucinations
Many fungi produce biologically active compounds, several of which
are toxic to animals or plants and are therefore called mycotoxins .
Of particular relevance to humans are mycotoxins produced by molds
causing food spoilage, and poisonous mushrooms (see above).
Particularly infamous are the lethal amatoxins in some Amanita
mushrooms, and ergot alkaloids , which have a long history of causing
serious epidemics of ergotism (St Anthony's Fire) in people consuming
rye or related cereals contaminated with sclerotia of the ergot
Claviceps purpurea . Other notable mycotoxins include the
aflatoxins , which are insidious liver toxins and highly carcinogenic
metabolites produced by certain
Aspergillus species often growing in
or on grains and nuts consumed by humans, ochratoxins , patulin , and
T-2 mycotoxin ) and fumonisins , which have
significant impact on human food supplies or animal livestock .
Mycotoxins are secondary metabolites (or natural products ), and
research has established the existence of biochemical pathways solely
for the purpose of producing mycotoxins and other natural products in
Mycotoxins may provide fitness benefits in terms of
physiological adaptation, competition with other microbes and fungi,
and protection from consumption (fungivory ). Many fungal secondary
metabolites (or derivatives) are used medically, as described under
Human Use below.
Ustilago maydis is a pathogenic plant fungus that causes smut disease
in maize and teosinte .
Plants have evolved efficient defense systems
against pathogenic microbes such as U. maydis. A rapid defense
reaction after pathogen attack is the oxidative burst where the plant
produces reactive oxygen species at the site of the attempted
invasion. U. maydis can respond to the oxidative burst with an
oxidative stress response, regulated by the gene
YAP1 . The response
protects U. maydis from the host defense, and is necessary for the
pathogen’s virulence. Furthermore, U. maydis has a well-established
DNA repair system which acts during mitosis and
meiosis. The system may assist the pathogen in surviving DNA damage
arising from the host plant’s oxidative defensive response to
Cryptococcus neoformans is an encapsulated yeast that can live in
both plants and animals. C. neoformans usually infects the lungs,
where it is phagocytosed by alveolar macrophages . Some C. neoformans
can survive inside macrophages, which appears to be the basis for
latency , disseminated disease, and resistance to antifungal agents.
One mechanism by which C. neoformans survives the hostile macrophage
environment is by up-regulating the expression of genes involved in
the oxidative stress response. Another mechanism involves meiosis .
The majority of C. neoformans are mating "type a". Filaments of mating
"type a" ordinarily have haploid nuclei, but they can become diploid
(perhaps by endoduplication or by stimulated nuclear fusion) to form
blastospores . The diploid nuclei of blastospores can undergo meiosis,
including recombination, to form haploid basidiospores that can be
dispersed. This process is referred to as monokaryotic fruiting. this
process requires a gene called
DMC1 , which is a conserved homologue
of genes recA in bacteria and
RAD51 in eukaryotes, that mediates
homologous chromosome pairing during meiosis and repair of DNA
double-strand breaks. Thus, C. neoformans can undergo a meiosis,
monokaryotic fruiting, that promotes recombinational repair in the
oxidative, DNA damaging environment of the host macrophage, and the
repair capability may contribute to its virulence.
Saccharomyces cerevisiae cells shown with DIC microscopy
The human use of fungi for food preparation or preservation and other
purposes is extensive and has a long history.
Mushroom farming and
mushroom gathering are large industries in many countries. The study
of the historical uses and sociological impact of fungi is known as
ethnomycology . Because of the capacity of this group to produce an
enormous range of natural products with antimicrobial or other
biological activities, many species have long been used or are being
developed for industrial production of antibiotics , vitamins, and
anti-cancer and cholesterol-lowering drugs. More recently, methods
have been developed for genetic engineering of fungi, enabling
metabolic engineering of fungal species. For example, genetic
modification of yeast species —which are easy to grow at fast rates
in large fermentation vessels—has opened up ways of pharmaceutical
production that are potentially more efficient than production by the
original source organisms.
Many species produce metabolites that are major sources of
pharmacologically active drugs. Particularly important are the
antibiotics, including the penicillins , a structurally related group
of β-lactam antibiotics that are synthesized from small peptides .
Although naturally occurring penicillins such as penicillin G
Penicillium chrysogenum ) have a relatively narrow
spectrum of biological activity, a wide range of other penicillins can
be produced by chemical modification of the natural penicillins.
Modern penicillins are semisynthetic compounds, obtained initially
from fermentation cultures, but then structurally altered for specific
desirable properties. Other antibiotics produced by fungi include:
ciclosporin , commonly used as an immunosuppressant during transplant
surgery ; and fusidic acid , used to help control infection from
methicillin-resistant Staphylococcus aureus bacteria. Widespread use
of antibiotics for the treatment of bacterial diseases, such as
tuberculosis , syphilis , leprosy , and others began in the early 20th
century and continues to date. In nature, antibiotics of fungal or
bacterial origin appear to play a dual role: at high concentrations
they act as chemical defense against competition with other
microorganisms in species-rich environments, such as the rhizosphere ,
and at low concentrations as quorum-sensing molecules for intra- or
interspecies signaling. Other drugs produced by fungi include
griseofulvin isolated from
Penicillium griseofulvum , used to treat
fungal infections, and statins (
HMG-CoA reductase inhibitors), used
to inhibit cholesterol synthesis . Examples of statins found in fungi
include mevastatin from
Penicillium citrinum and lovastatin from
Aspergillus terreus and the oyster mushroom .
Traditional And Folk Medicine
The medicinal fungi
Ganoderma lucidum (left) and
Ophiocordyceps sinensis (right)
Certain mushrooms enjoy usage as therapeutics in folk medicines ,
Traditional Chinese medicine . Notable medicinal mushrooms
with a well-documented history of use include
Agaricus subrufescens ,
Ganoderma lucidum , and
Ophiocordyceps sinensis . Research has
identified compounds produced by these and other fungi that have
inhibitory biological effects against viruses and cancer cells .
Specific metabolites, such as polysaccharide-K , ergotamine , and
β-lactam antibiotics , are routinely used in clinical medicine. The
shiitake mushroom is a source of lentinan , a clinical drug approved
for use in cancer treatments in several countries, including
Japan , polysaccharide-K (brand name Krestin), a
chemical derived from
Trametes versicolor , is an approved adjuvant
for cancer therapy.
Baker\'s yeast or
Saccharomyces cerevisiae , a unicellular fungus, is
used to make bread and other wheat-based products, such as pizza dough
and dumplings .
Yeast species of the genus
Saccharomyces are also
used to produce alcoholic beverages through fermentation.
Aspergillus oryzae ) is an essential ingredient in brewing Shoyu
(soy sauce ) and sake , and the preparation of miso , while Rhizopus
species are used for making tempeh . Several of these fungi are
domesticated species that were bred or selected according to their
capacity to ferment food without producing harmful mycotoxins (see
below), which are produced by very closely related Aspergilli . Quorn
, a meat substitute , is made from
Fusarium venenatum .
EDIBLE AND POISONOUS SPECIES
Amanita phalloides accounts for the majority of fatal mushroom
Edible mushrooms are well-known examples of fungi. Many are
commercially raised, but others must be harvested from the wild.
Agaricus bisporus , sold as button mushrooms when small or Portobello
mushrooms when larger, is a commonly eaten species, used in salads,
soups, and many other dishes. Many Asian fungi are commercially grown
and have increased in popularity in the West. They are often available
fresh in grocery stores and markets, including straw mushrooms
Volvariella volvacea ), oyster mushrooms (
Pleurotus ostreatus ),
Lentinula edodes ), and enokitake (
There are many more mushroom species that are harvested from the wild
for personal consumption or commercial sale. Milk mushrooms , morels ,
chanterelles , truffles , black trumpets , and porcini mushrooms
Boletus edulis ) (also known as king boletes) demand a high price on
the market. They are often used in gourmet dishes.
Certain types of cheeses require inoculation of milk curds with
fungal species that impart a unique flavor and texture to the cheese.
Examples include the blue color in cheeses such as Stilton or
Roquefort , which are made by inoculation with
. Molds used in cheese production are non-toxic and are thus safe for
human consumption; however, mycotoxins (e.g., aflatoxins, roquefortine
C , patulin, or others) may accumulate because of growth of other
fungi during cheese ripening or storage.
Stilton cheese veined
Many mushroom species are poisonous to humans, with toxicities
ranging from slight digestive problems or allergic reactions as well
as hallucinations to severe organ failures and death. Genera with
mushrooms containing deadly toxins include
Lepiota , and, the most infamous,
Amanita . The latter genus includes
the destroying angel (A. virosa ) and the death cap (A. phalloides ),
the most common cause of deadly mushroom poisoning. The false morel
Gyromitra esculenta ) is occasionally considered a delicacy when
cooked, yet can be highly toxic when eaten raw. Tricholoma equestre
was considered edible until it was implicated in serious poisonings
causing rhabdomyolysis . Fly agaric mushrooms (
Amanita muscaria) also
cause occasional non-fatal poisonings, mostly as a result of ingestion
for its hallucinogenic properties. Historically, fly agaric was used
by different peoples in
Europe and Asia and its present usage for
religious or shamanic purposes is reported from some ethnic groups
such as the Koryak people of north-eastern
As it is difficult to accurately identify a safe mushroom without
proper training and knowledge, it is often advised to assume that a
wild mushroom is poisonous and not to consume it.
Grasshoppers killed by
In agriculture, fungi may be useful if they actively compete for
nutrients and space with pathogenic microorganisms such as bacteria or
other fungi via the competitive exclusion principle , or if they are
parasites of these pathogens. For example, certain species may be used
to eliminate or suppress the growth of harmful plant pathogens, such
as insects, mites , weeds , nematodes , and other fungi that cause
diseases of important crop plants. This has generated strong interest
in practical applications that use these fungi in the biological
control of these agricultural pests.
Entomopathogenic fungi can be
used as biopesticides , as they actively kill insects. Examples that
have been used as biological insecticides are
Beauveria bassiana ,
Paecilomyces (Isaria) spp, and
Lecanicillium lecanii .
Endophytic fungi of grasses of the genus
Neotyphodium , such as N. coenophialum , produce alkaloids that are
toxic to a range of invertebrate and vertebrate herbivores . These
alkaloids protect grass plants from herbivory , but several endophyte
alkaloids can poison grazing animals, such as cattle and sheep.
Infecting cultivars of pasture or forage grasses with Neotyphodium
endophytes is one approach being used in grass breeding programs; the
fungal strains are selected for producing only alkaloids that increase
resistance to herbivores such as insects, while being non-toxic to
Certain fungi, in particular "white rot" fungi, can degrade
insecticides , herbicides , pentachlorophenol , creosote , coal tars ,
and heavy fuels and turn them into carbon dioxide , water, and basic
elements. Fungi have been shown to biomineralize uranium oxides ,
suggesting they may have application in the bioremediation of
radioactively polluted sites.
Several pivotal discoveries in biology were made by researchers using
fungi as model organisms , that is, fungi that grow and sexually
reproduce rapidly in the laboratory. For example, the one gene-one
enzyme hypothesis was formulated by scientists using the bread mold
Neurospora crassa to test their biochemical theories. Other important
model fungi are
Aspergillus nidulans and the yeasts Saccharomyces
Schizosaccharomyces pombe , each of which with a long
history of use to investigate issues in eukaryotic cell biology and
genetics , such as cell cycle regulation, chromatin structure, and
gene regulation . Other fungal models have more recently emerged that
address specific biological questions relevant to medicine , plant
pathology , and industrial uses; examples include
Candida albicans , a
dimorphic, opportunistic human pathogen,
Magnaporthe grisea , a plant
Pichia pastoris , a yeast widely used for eukaryotic
protein production .
Fungi are used extensively to produce industrial chemicals like
citric , gluconic , lactic , and malic acids, and industrial enzymes,
such as lipases used in biological detergents , cellulases used in
making cellulosic ethanol and stonewashed jeans , and amylases ,
invertases , proteases and xylanases . Several species, most notably
Psilocybin mushrooms (colloquially known as magic mushrooms), are
ingested for their psychedelic properties, both recreationally and
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